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However, when many previously proposed evolutionary novelties have been illuminated by genetic, developmental, and fossil data, they have refined and narrowed our concept of biological \u201cnewness.\u201d For example, they show that these novelties can occur at one or multiple levels of biological organization. Here, we review the identity of structures in the avian vocal organ, the syrinx, and bring together developmental data on airway patterning, structural data from across tetrapods, and mathematical modeling to assess what is novel. In contrast with laryngeal cartilages that support vocal folds in other vertebrates, we find no evidence that individual cartilage rings anchoring vocal folds in the syrinx have homology with any specific elements in outgroups. Further, unlike all other vertebrate vocal organs, the syrinx is not derived from a known valve precursor, and its origin involves a transition from an evolutionary \u201cspandrel\u201d in the respiratory tract, the site where the trachea meets the bronchi, to a target for novel selective regimes. We find that the syrinx falls into an unusual category of novel structures: those having significant functional overlap with the structures they replace. The syrinx, along with other evolutionary novelties in sensory and signaling modalities, may more commonly involve structural changes that contribute to or modify an existing function rather than those that enable new functions.<\/jats:p>","DOI":"10.1073\/pnas.1804586115","type":"journal-article","created":{"date-parts":[[2018,9,24]],"date-time":"2018-09-24T15:04:26Z","timestamp":1537801466000},"page":"10209-10217","update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":58,"title":["Identity and novelty in the avian syrinx"],"prefix":"10.1073","volume":"115","author":[{"given":"Evan P.","family":"Kingsley","sequence":"first","affiliation":[{"name":"Department of Genetics, Harvard Medical School, Boston, MA 02115;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8426-0373","authenticated-orcid":false,"given":"Chad M.","family":"Eliason","sequence":"additional","affiliation":[{"name":"Integrative Research Center, Field Museum of Natural History, Chicago, IL 60605;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tobias","family":"Riede","sequence":"additional","affiliation":[{"name":"Department of Physiology, Midwestern University, Glendale, AZ 85308;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhiheng","family":"Li","sequence":"additional","affiliation":[{"name":"Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences Center for Excellence in Life and Paleoenvironment, Chinese Academy of Sciences 100044 Beijing, China;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tom W.","family":"Hiscock","sequence":"additional","affiliation":[{"name":"Li Ka Shing Centre, Cambridge Institute, Cancer Research UK, Cambridge CB2 0RE, United Kingdom;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Farnsworth","sequence":"additional","affiliation":[{"name":"Department of Mechanical Engineering, Brigham Young University, Provo, UT 84602;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Scott L.","family":"Thomson","sequence":"additional","affiliation":[{"name":"Department of Mechanical Engineering, Brigham Young University, Provo, UT 84602;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Franz","family":"Goller","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Utah, Salt Lake City, UT 84112;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Clifford J.","family":"Tabin","sequence":"additional","affiliation":[{"name":"Department of Genetics, Harvard Medical School, Boston, MA 02115;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julia A.","family":"Clarke","sequence":"additional","affiliation":[{"name":"Department of Geological Sciences, University of Texas at Austin, Austin, TX 78712"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2018,9,24]]},"reference":[{"key":"e_1_3_3_1_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.1253293"},{"key":"e_1_3_3_2_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.es.22.110191.001305"},{"key":"e_1_3_3_3_2","doi-asserted-by":"publisher","DOI":"10.1038\/41710"},{"key":"e_1_3_3_4_2","doi-asserted-by":"publisher","DOI":"10.1002\/jez.b.21425"},{"key":"e_1_3_3_5_2","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.117101"},{"key":"e_1_3_3_6_2","doi-asserted-by":"publisher","DOI":"10.1002\/1097-4687(200009)245:3<241::AID-JMOR5>3.0.CO;2-H"},{"key":"e_1_3_3_7_2","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10236"},{"key":"e_1_3_3_8_2","doi-asserted-by":"publisher","DOI":"10.1038\/nature19852"},{"key":"e_1_3_3_9_2","doi-asserted-by":"publisher","DOI":"10.1080\/08912960903033327"},{"key":"e_1_3_3_10_2","doi-asserted-by":"crossref","unstructured":"KC Colafrancesco M Gridi-Papp Vocal sound production and acoustic communication in amphibians and reptiles. 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